Patterns of Male Production in the Stingless Bee Melipona Favosa (Apidae, Meliponini) Tong Chinhinst, Gijs Grob, Francis Meeuwsen, Marinus Sommeijer
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Patterns of male production in the stingless bee Melipona favosa (Apidae, Meliponini) Tong Chinhinst, Gijs Grob, Francis Meeuwsen, Marinus Sommeijer To cite this version: Tong Chinhinst, Gijs Grob, Francis Meeuwsen, Marinus Sommeijer. Patterns of male production in the stingless bee Melipona favosa (Apidae, Meliponini). Apidologie, Springer Verlag, 2003, 34 (2), pp.161-170. 10.1051/apido:2003008. hal-00891768 HAL Id: hal-00891768 https://hal.archives-ouvertes.fr/hal-00891768 Submitted on 1 Jan 2003 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Apidologie 34 (2003) 161–170 © INRA/DIB-AGIB/ EDP Sciences, 2003 161 DOI: 10.1051/apido:2003008 Original article Patterns of male production in the stingless bee Melipona favosa (Apidae, Meliponini) Tong X. CHINHa,b*, Gijs B.J. GROBc, Francis J.A.J. MEEUWSENa, Marinus J. SOMMEIJERa a Department of Social Insects, Utrecht University, PO Box 80.086, 3508 TB Utrecht, The Netherlands b National Bee Research and Development Centre, Langha, Dongda, Hanoi, Vietnam c Nationaal Herbarium Nederland, Leiden University, PO Box 9514, 2300 RA Leiden, The Netherlands (Received 4 April 2002; revised 3 August 2002; accepted 17 September 2002) Abstract – In many stingless bee species, laying workers oviposit trophic eggs that serve as a component of the queen’s diet. Workers of some species also lay reproductive worker eggs that give rise to males. Male- producing workers can occur in queenright colonies. We studied male production by workers of Melipona favosa. In six colonies monitored under field conditions, we observed that males emerged during distinct “Male Emerging Periods”. Subsequently, we studied the laying of male eggs in laboratory observation hives with the use of video. We found that laying workers oviposited reproductive eggs in distinct “Reproductive Laying Worker Periods” and that this was followed by the clumped mergence of males afterwards. Behaviours of laying workers and of the ovipositing queen are preliminarily described. Melipona / laying worker / male production / reproductive eggs / trophic eggs 1. INTRODUCTION sons than to other worker’s sons (Ratnieks, 1988, Peters et al., 1999). The haplo-diploid system in social In colonies of the honey bee, Apis mellifera, Hymenoptera influences the relatedness of which are headed by a highly polyandrous queens and workers to males and can queen, males are normally produced only by contribute to conflict over male production. In the queen (Seeley, 1985; Peters et al., 1999). the light of the kin selection theory (Hamilton, In queenless colonies, honeybee workers can 1972), when a colony is headed by a single activate their ovaries and lay unfertilised eggs. monandrous queen, workers are expected to Workers in queenright colonies generally do favour their sons (relatedness, r = 0.5) or other not activate their ovaries as a result of worker’s sons (nephews) (r = 0.375) above pheromonal control by the queen (Velthuis, sons of the queen (brothers) (r = 0.25) 1970). However, it has recently been (Hamilton, 1964; Ratnieks, 1988, Peters et al., demonstrated that queen-right Apis mellifera 1999). In contrast, in colonies headed by workers sometimes do lay eggs but that their polyandrous queens, worker-queen conflict contribution to male production is low because over male production may be reduced because of worker policing (Barron et al., 2001; workers are then more related to the queen’s Ratnieks,1993; Visscher, 1989). * Correspondence and reprints E-mail: [email protected] 162 T.X. Chinh et al. In contrast to Apis, stingless bee both the queen and the workers, and try to (Meliponini) workers with active ovaries are explain the observed patterns in light of exist- common in queenright colonies (Sakagami, ing theories. 1982). Moreover, worker oviposition in queenright colonies occurs widely among the various groups of stingless bees (Sakagami, 2. MATERIALS AND METHODS 1982; Sommeijer and van Buren, 1992; Imperatriz-Fonseca and Kleinert, 1998; All colonies of M. favosa were kept in wooden Koedam et al., 1999). However, in queenright box hives (W × L × H: 20.5 × 35.5 × 15 cm), which colonies, Melipona workers may lay mostly were covered by glass lids facilitating behavioural trophic, not reproductive eggs (Sakagami, observation. The mature brood combs were 1982; Sommeijer et al., 1984a). Trophic eggs partially separated from the major brood-nest by are eaten by the queen prior to her oviposition, placing them in small annex boxes that were while reproductive eggs can give rise to off- connected to the major nest box with plastic tubes. spring. The annex served as a place where newly emerged bees could remain for about the first 24 hours after In some species of non-Melipona stingless their emergence on the comb. Recently eclosed bees, e.g., in Scaptotrigona, reproductive workers and males of M. favosa do not disperse workers commonly oviposit after queen ovi- from the comb of emergence. Upon the recording of position (Beig, 1972; Bego, 1982; Sakagami, daily-emerged adults, these were reintroduced into 1982), which leads to cells containing both a the main brood-nest compartment. Colony worker egg and a queen egg. In contrast, in development parameters such as daily cell Melipona, oviposition by reproductive laying production, numbers of new cells and storage pots, workers generally precludes oviposition by the etc., were measured weekly. queen in the same cell (Sommeijer et al., The field studies were carried out from October 1984b; Sommeijer and van Buren, 1992). 1994 to February 1995 on the island of Tobago Reproductive workers of M. favosa may also (Trinidad and Tobago, West Indies). Six queenright colonies of Melipona favosa that earlier had been oviposit after queen oviposition, however this installed in a room in a wooden building were used phenomenon is very rare. to study the temporal patterns of male production. Melipona trophic and reproductive worker These colonies had free flight to the outside in their eggs differ in their external morphology. natural habitat through plastic tubes leading Reproductive eggs in queenright colonies through the wall. The colonies received no resemble reproductive worker eggs layed additional feeding. The colonies had been in this in queenless colonies (van Buren and position for more than two years. The population of these colonies ranged from 129 to 204 bees, which Sommeijer, 1988), and have a well-developed is normal for this species. chorion. In contrast, the chorion of trophic eggs is poorly developed. However, these In 1997, four queenright colonies (referred to as 7, 8, 9 and 10) of M. favosa were housed in the room differences are graded and not discrete of our laboratory at the Faculty of Biology, Utrecht (Sommeijer et al., 1984a). Besides a difference University, The Netherlands. These colonies in trophic and reproductive egg morphology, originated from Trinidad and Tobago and their there is a difference in the behaviour of populations ranged from 109 to 270 bees1 workers that lay trophic and reproductive Temperature was kept constantly at 24 degrees eggs, so the two categories of laying workers Celsius and RH was constantly around 75%. In can be easily distinguished (Sommeijer and these lab colonies, all ovipositions were analysed to van Buren, 1992). identify whether the queen or a worker oviposited in a cell. Red light and video were used to observe We have studied the production of males in the ovipositions, comb development and adult M. favosa. In a previous paper we showed that emergence continuously and simultaneously from in M. favosa the workers are the main produc- April to November. To identify the maternity of the ers of males (Sommeijer et al., 1999). In this emerged bees, the location of each oviposited cell paper we describe the production of males in more detail and show that it occurs periodi- 1 The colonies studied in the Utrecht lab had an cally both under field and laboratory condi- average population of 194 and were larger than the tions. In the discussion we link this clumped colonies studied in the field (average population 161), male production to oviposition behaviour of but this difference was not significant (P > 0.05). Observations on male production in Melipona favosa 163 Table I. Some parameters of all the fifteen observed Male Emerging Periods in the field and in the lab. MEP Colony Start End Duration Males % Males Intervals Abs. max Rel. max (day) (day) 1 1 11 Nov 3 Jan 53 153 14.6 9 1 Dec 10 Dec 2 1 24 Jan 23 Feb 30 92 13.4 2 4 Feb 8 Feb 3 2 24 Nov 8 Dec 14 8 11.0 1 1 Feb 1 Dec 4 2 17 Jan 23 Feb 37 62 15.3 1 27 Jan 26 Jan 5 3 19 Dec 15 Feb 74 88 18.1 6 17 Jan 17 Jan 6 4 15 Nov 19 Dec 35 19 9.7 5 26 Dec 18 Dec 7 4 24 Jan 23 Feb 30 79 16.0 2 14 Feb 1 Feb 8 5 31 Oct 17 Nov 17 43 23.2 1 11 Dec 14 Nov 9 5 29 Dec 22 Jan 24 12 3.6 4 13 Jan 4 Jan 10 5 11 Feb 21 Feb 10 14 14.9 1 17 Feb 11 Feb 11 6 16 Dec 22 Jan 37 69 37.3 2 3 Jan 2 Jan 12 7* 12 May 27 Jun 46 422 70.7 0 25 May 17, 19, 24 May 13 8* 8 Sep 17 Oct 38 163 38.0 1 21 Sep 29, 23 Sep 14 9* 10 Apr 16 Jun 67 940 10.0 3 21 Apr 23, 24 Apr; 3 Jun 15 10* 9 Sep 7 Dec 60 192 33.8 5 10 Oct 10, 16, 17 Oct Average 38.1 157.1 22.0 2.9 St.dev.